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Dapedium Sp. from the Toarcian (Lower Jurassic) Úrkút Manganese Ore Formation (Bakony Mts., Hungary) and an Overview of Diversity of Dapediiform Fishes

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Dapedium Sp. from the Toarcian (Lower Jurassic) Úrkút Manganese Ore Formation (Bakony Mts., Hungary) and an Overview of Diversity of Dapediiform Fishes Palaeobiodiversity and Palaeoenvironments https://doi.org/10.1007/s12549-019-00390-7 ORIGINAL PAPER Dapedium sp. from the Toarcian (Lower Jurassic) Úrkút Manganese Ore Formation (Bakony Mts., Hungary) and an overview of diversity of dapediiform fishes Márton Szabó1,2 & József Pálfy3,4 Received: 18 August 2018 /Revised: 10 November 2018 /Accepted: 28 May 2019 # The Author(s) 2019 Abstract Dapediidae are a characteristic group of deep-bodied Mesozoic actinopterygian fishes with a moderate diversity at genus- and species- level. Here, we add a new occurrence to their patchy fossil record and describe in detail a nearly complete dapediid specimen from the pelagic deposit of the Toarcian Úrkút Manganese Ore Formation in the Transdanubian Range of Hungary. The preserved characters represent nearly all anatomically important body parts and allow assignment to Dapedium and comparison with other dapediid genera. This is the first reported occurrence of the order in Hungary and the Carpathian Basin that extends the known geographical range of the genus to the Mediterranean (western Tethyan) Jurassic. A review of the temporal distribution of published occurrences of dapediids permits speculation that the disappearance of exclusively Late Triassic genera, coincident with the end-Triassic extinction event, was likely related to their specialised feeding strategies and light or incomplete squamation. Multiple environmental crises (warming, acidification and anoxia) severely affected reefal habitats and favoured the survival of the generalist-durophagous Dapedium.The Úrkút specimen adds important data to the Early Jurassic, particularly Toarcian Lagerstätte-dominated fossil record of dapediids. In contrast to the end-Triassic, the Toarcian oceanic anoxic event did not lead to genus extinction among dapediids, possibly prevented by adaptations evolved during the preceding and similarly multi-stressor event. Continuing studies of Mesozoic fish specimens in Hungarian collections may provide new records and insights into other groups as well. Keywords Dapedium . Toarcian . End-Triassic . Extinction . Diversity Introduction interlocking rhomboidal ganoid scales, relatively small mouth, skull bones ornamented by ganoine-tubercles and ridges, hem- Dapedium Leach, 1822 is an extinct genus of lower shaped dorsal and anal fins, dorsal fins with more than 20 rays, actinopterygian fish, first discovered at Lyme Regis, England. It and pronounced dorsal and ventral ridge scales (Thies and Hauff is characterised by a deep (hypsisomatic), disc-like body, 2011; Smithwick 2015; Thies and Waschkewitz 2015;Gibson 2016). The fossil record of Dapedium ranges from the late Norian (Late Triassic) (Tintori 1983; Lombardo and Tintori 2005)tothe * Márton Szabó earliest Aalenian (Middle Jurassic) (Thies and Hauff 2011; [email protected] Maxwell and López-Arbarello 2018). All confirmed occurrences have been reported from Europe; the only alleged Dapedium find József Pálfy from India (Jain 1973) has been disputed (Maxwell and López- [email protected] Arbarello 2018). The majority of the material originates from well-known Lagerstätten and strata with exceptional preservation 1 Department of Paleontology, Eötvös University, Pázmány Péter sétány 1/C, Budapest 1117, Hungary in England, France, Belgium, Italy and Germany (i.e. sites in the western Tethyan shelf or adjoining epicontinental seas). This list 2 Department of Palaeontology and Geology, Hungarian Natural History Museum, Ludovika tér 2, Budapest 1083, Hungary is augmented here with a find from a Toarcian black shale-hosted manganiferous ore deposit in Hungary, representing a relatively 3 Department of Geology, Eötvös University, Pázmány Péter sétány 1/C, Budapest 1117, Hungary deep, pelagic environmental setting. On the basis of tooth morphology and functional mor- 4 MTA-MTM-ELTE Research Group for Paleontology, Ludovika tér 2, Budapest 1083, Hungary phology of the jaw, Dapedium was interpreted as a Palaeobio Palaeoenv generalist well-adapted for generalist durophagy (Thies and Geological Survey of Hungary (MGSH) in the same condition Hauff 2011;Smithwick2015;Stumpfetal.2017). as originally illustrated. Főzy and Szente (2014, p. 170) figured Facultative durophagy may have helped Dapedium in times an additional specimen, which is also housed in the vertebrate of increased competition for food resources (Smithwick fossil collection of the MGSH. Mineralogical and geochemical 2015). Based on a spectacular find of a Dapedium stuck studies of the phosphoritic nodules that yielded the fish remains in a shell of a Lytoceras ammonite, scavenging mode of life from Úrkút were carried out by Polgári et al. (2004). was also suggested (Thies and Hauff 2011,fig.13B). The Úrkút Manganese Ore Formation that yielded the According to Thies and Hauff (2012), Dapedium was also dapediid specimen described here is a hitherto less known a prey animal for larger predators. source of exceptional preservation related to bottom water The order Dapediiformes (including only the family anoxia. The relation of the Úrkút manganese ore deposit to Dapediidae) was established by Thies and Waschkewitz (2015), the Toarcian Oceanic Anoxic Event (T-OAE or Jenkyns event, who separated dapediids from semionotiform fishes, to which Müller et al. 2017) is well-established (see overviews by Haas they were formerly attributed. However, the phylogenetic study 2012 and Polgári et al. 2016a, b). The biological productivity was restricted to Dapedium. Within Dapediidae, Thies and Hauff was high in the surface waters, but bottom waters were (2011) included seven genera (Hemicalypterus, Tetragonolepis, oxygen-deficient (Polgári et al. 1991). Paradapedium, Heterostrophus, Dapedium, Dandya and Sargodon). Gibson (2016) referred an additional genus, Aetheolepis, to the family. In 2018, a ninth genus Scopulipiscis Geological setting was introduced and described by Latimer and Giles (2018). Since the late 1960s, several fish fossils have been reported The village of Úrkút is located in Veszprém County, in the south- from the Úrkút Magnanese Ore Formation in Hungary (see in ern Bakony Mountains, western Hungary (Fig. 1a). Manganese “Previous work” section). A few of these remains are now ore mining in the Bakony Mountains started here in the Úrkút housed in museum collections, including one specimen bearing Basin in 1917 and was terminated in 2016. The black shale- diagnostic characters of Dapediidae. The aim of the present study hosted Mn-oxide and carbonate ore body of Úrkút is among is to provide a detailed anatomical and taxonomical description the ten largest deposits in the world (Polgári et al. 2013). of this specimen. A summary of the available information allows The Úrkút Manganese Ore Formation occurs in a large area in us to consider the history of diversification of the group, with an the Úrkút Basin. Its thickness reaches 30 m in the western part of emphasis on the possible reasons for selective extinction and the basin, and exceeds 100 m in the north-western part of the survival of dapediiforms at the end-Triassic and Toarcian events. basin, it includes three ore beds (Polgári et al. 1991;Bíró2014). The Úrkút ore deposits occur in a NE-SW trending, ~ 12-km- long and 4- to 6-km-wide zone. The manganiferous ore conform- Previous work ably overlies the Pliensbachian cherty limestone of the Isztimér Formation (Polgári et al. 2012, 2013)(Fig.1b). The sequence Mesozoic fish remains from Hungary have been the subject of starts with ~ 0.5 m of Mn-oxide ore, overlain by 1.0 m of radi- only a few studies. Toarcian fishes, representing several spe- olarian clay marlstone. The main manganese ore body has a cies, are figured from the black shale facies exposed in the maximum thickness of 15 m and is made up of variously Réka Valley (Mecsek Mountains, southern Hungary) (Főzy coloured and finely laminated Mn ore. This unit is followed by and Szente 2014), but have not yet been studied in detail. ~ 10 m of radolarian clay marlstone and ~ 2.5 m of Dulaietal.(1992) identified only a single species, manganiferous clay marlstone. The uppermost part of the se- Leptolepis normandica. With the exception of the Réka quence consists of 2.5 m of radiolarian clay marlstone, and ~ Valley material, all other reports are based on isolated remains. 2.5 m of manganiferous clay marlstone. A discontinuous chert Although fossil fishes of the Úrkút Manganese Ore Formation layer (Cservár Flintstone Member) represents the closes part of have been known for more than 50 years, published information the formation (Bíró 2014). For further details on the geology and is restricted to only a few brief reports. A large specimen was stratigraphy of the Úrkút Mn ore deposit, see Szabó and found and figured by Cseh-Németh (1966)as“ganoidfishre- Grasselly (1980), Szabó et al. (1981), Grasselly and Pantó main” (note that this specimen is the subject of the present study). (1988),Polgárietal.(1991, 2012, 2013, 2016a, b), Pantó et al. Polgári et al. ( 2000)andPászti(2004) also figured this specimen ( 1996), Haas (2012)andBíró(2014). as a pycnodontid fish fossil. Pászti (2004) gave the first detailed account on the fish remains of the Úrkút Manganese Ore Formation, describing them as pycnodontiform and amiiform Material and methods fishes. However, from the three specimens figured by Pászti (2004), only Cseh-Németh’s(1966) specimen was found still Of the four Toarcian fossil fish specimens known from Úrkút, available in the vertebrate fossil collection of the Mining and the present study is focused
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